ASIAN MYRMECOLOGY Volume 7, 1 – 13, 2015

ISSN 1985-1944 © Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat,
Pudjianto, Dadan Hindayana, Yann Clough and Teja Tscharntke, Damayanti Buchori

Agricultural land use alters species composition

but not species richness of ant communities

Ratna Rubiana1*, Akhmad Rizali2, Lisa H. Denmead3,

Winda Alamsari1, Purnama Hidayat1, Pudjianto1, Dadan Hindayana1,
Yann Clough3 and Teja Tscharntke3, Damayanti Buchori1
1
Department of Plant Protection, Faculty of Agriculture, Bogor Agricultural
University. Jl. Kamper, Kampus IPB Dramaga, Bogor, 16680 Indonesia
2
Department of Plant Pests and Diseases, Faculty of Agriculture, University
of Brawijaya. Jl. Veteran, Malang, 65145 Indonesia
3
Agroecology, Georg-August-University, Grisebachstr. 6, 37077,
Göttingen, Germany

*Corresponding author’s email: ratna.rubiana@gmail.com

ABSTRACT. Land-use change causes undesirable effects such as biodiversity

decline, altered community structure and reduced ecosystem services. Changes
in species composition and disrupted trophic interactions between pests and
their natural enemies may also result causing decreased ecosystem services. We
studied the effects of forest habitat transformation on the community structure
of ants, which include major biological control agents. We focused on four
types of land use around Harapan Forest (Harapan) and Bukit Duabelas Nation-
al Park (BDNP), Jambi, Sumatra, Indonesia: forest, jungle rubber, rubber plan-
tations and oil palm plantations. Four replicate patches of each land-use type
were sampled, with plot sizes of 50 x 50 m at each of the 32 sites. Ants were
collected by hand in combination with tuna and sugar baiting on three strata i.e.
leaf litter, soil and tree. We found 104 ant species in total. Surprisingly, ant spe-
cies richness per plot was not significantly different among land-use types, both
in Harapan and BDNP. However, few ant species were shared among different
land-use types. Forest and jungle rubber communities are relatively similar to
each other (but still different), and distinct from communities in oil palm and
rubber plantations. We conclude that conversion of remnant forested habitats
to plantations would result in a net loss of ant species, even though ant species
richness in plantations and forested habitats are similar.

Keywords: land-use change, hand-collecting, oil palm, rubber, Sumatra island

INTRODUCTION vival of many organisms (Pringle 2007). When

Habitat transformation is an unfortunate conse-
quence of human population increase. Natural
habitats ever-growing are being altered by an-
thropogenic activities (Morris 2010). Habitat
transformation degrades natural habitats and in-
terferes with the resources necessary for the sur-
their habitat is destroyed, plants and animals that
had occupied the habitat are often displaced or
destroyed, thus reducing biodiversity and en-
hancing the likelihood of extinction (Swift et al.
2004). Therefore, habitat transformation is one
of the major causes of biodiversity decline along
with climate change, nitrogen deposition and in-

11 - AM7 - Agricultural land use alters species composition.indd 1 16-Jul-15 3:55:38 PM

 2 Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat, Pudjianto,
Dadan Hindayana, Yann Clough & Teja Tscharntke, Damayanti Buchori
creased atmospheric CO2 concentration (Sala et
al. 2000).
Biodiversity is important in regulating
and sustaining the direct and indirect contribu-
tions of ecosystems to human (ecosystem ser-
vices) (Alberti 2005). The reduction of species
richness often causes decreases in ecosystem ser-
vices (Naeem et al. 1999). In agricultural produc-
tion systems, insects provide ecosystem services
such as pest control, pollination, and soil fertility
(Power 2010). Decreasing the number of species
in economically important functional groups may
lead to increased pest density, reduced pollinator
and natural enemies services (Tscharntke et al.
2012). Ants (Hymenoptera: Formicidae) provide
important ecosystem services including biologi-
cal pest control, seed dispersal, and soil modifi-
cation (Hill & Hoy 2003; Gammans et al. 2005;
Lach et al. 2010; Philpott et al. 2010). However,
ants are sensitive to changes in their environment
including changes in dominant vegetation struc-
ture, food availability, and nesting resources (An-
dersen 2000). The changes of vegetation struc-
ture resulting from forest transformation usually
experience changes in ant community structure
(Nakamura et al. 2007). Habitat transformation
may severely impact the abundance, community
structure, and interaction behavior of ants toward
each other and other organisms (e.g. avoidance of
predators and parasitism) (Kaspari et al. 2003).
Due to the benefits of ants for ecosystem services
(Wielgoss et al. 2013), as well as their sensitivity
to change, they are an ideal focus group to inves-
tigate the impacts of habitat transformation.
Here, we compare ant communities in
remnant forested habitats of Jambi province, Su-
matra, with those found in several common agri-
cultural land-use types: rubber agroforests with
diverse vegetation (jungle rubber), monoculture
rubber and oil palm plantations. The objectives
of this research were to (1) compare the diver-
sity of ants in the different types of land use, (2)
compare the species composition and community
structure across the different habitat types, and
(3) investigate changes in ant dominance patterns
resulting from transformation of their habitat.
11 - AM7 - Agricultural land use alters species composition.indd 2
MATERIALS AND METHODS
Study sites
Fieldwork was conducted in the tropical lowland
rainforest in Jambi Province in southwest Suma-
tra, Indonesia (Fig. 1). Two sites were chosen
for this research: Bukit Duabelas National Park
(BDNP) and Harapan Forest (Harapan). The hab-
itat transformation systems investigated consisted
of lowland rainforest, jungle rubber (extensively
managed rubber plantations, which have been
logged at least once, but usually more often), and
intensive rubber and oil palm plantations. In each
of the two areas, four sites (plot size 50 x 50 m)
in each type of land use were established, for a to-
tal of 32 study plots. Each plot had five sub-plots
(5 x 5 m) defined for sample collection. Subplot
location was determined randomly, and was reas-
signed for every plot.
Sample collection and identification
We used both direct sampling and baiting of ants.
Direct sampling allowed estimation of the num-
ber of ant species per unit area. Direct sampling
in each stratum (leaf litter, soil, and tree) lasted 5
- 10 min. Leaf litter was separated into coarse and
fine litter and ants were taken from the fine litter
in the tray. For the soil strata, ants were collected
directly from the ground with forceps. Sampling
on trees was combined with baiting, using tuna
and sugar bait to attract the ants (Bestelmeyer et
al. 2000). Sugar water and canned tuna were put
in a plastic plate with a diameter of 20 cm with
4 bait containers with a diameter of 2 cm. Sugar
water was absorbed into a foam that was placed
in the container. Baits were installed for 1 hour.
Ant sampling was completed between 09.00 and
11.00 am from 22 February to 31 March 2013 and
only carried out during sunny weather.
All specimens were stored in 70% etha-
nol and were identified to morphospecies using a
stereo microscope and an identification guide for
Bornean Ants (Hashimoto 2003).
Data Analysis
To understand whether ant species richness dif-
fered between habitat types, we used an analysis
16-Jul-15 3:55:38 PM
 Agricultural land use alters species composition but not species richness of ant communities 3

Table 1. Ant species richness in four land-use types in Bukit Duabelas National Park (BDNP) and Harapan Forest.
The difference of ant species richness between land-use types on each site was tested using ANOVA.

Land-use Subfamily Genus Species Average Statistic

BDNP
Primary forest 5 27 42 17.5
F3,10 = 1.26
Jungle rubber 5 22 31 14.0 P = 0.340
Rubber plantation 5 29 45 21.5
Oil palm plantation 5 27 40 21.3
Sub total 6 50 86 39.5
Harapan Forest
Primary forest 5 26 42 19.3
F3,15 = 0.37
Jungle rubber 5 29 48 19.5 P = 0.779
Rubber plantation 5 25 45 20.5
Oil palm plantation 5 25 43 17.8
Sub total 5 38 81 44.5
Total 6 52 104

Table 2. Dissimilarity of ant species (Bray-Curtis index) between different land-use types in Bukit Duabelas and
Harapan sites. The first letter indicates landscape (B: Bukit Duabelas, H: Harapan) and the second letter indicates
the land-use type (F: forest, J: jungle, O: Oil palm, R: rubber)

Land-use BF BJ BO BR HF HJ HO HR
BF 0
BJ 0.45 0
BO 0.61 0.61 0
BR 0.54 0.53 0.48 0
HF 0.36 0.36 0.51 0.49 0
HJ 0.52 0.53 0.53 0.47 0.45 0
HO 0.53 0.49 0.37 0.39 0.58 0.50 0
HR 0
0.56 0.47 0.43 0.27 0.42 0.44 0.30

of variance (ANOVA). Ant community structure RESULTS

was compared between different land-use types
within each study area based on Bray-Curtis dis-
similarity index and further analyzed using non-
metric multidimensional scaling (NMDS). Sig-
nificance tests for differences in community com-
position between land-use types were performed
using the analysis of similarity test (ANOSIM;
Clarke 1993). All analyses were performed using
R statistic (R Core Team 2014).
A total of 104 ant species were collected, repre-
senting six subfamilies and 52 genera (Table 1).
Species richness in the BDNP site (86 species)
was slightly higher than in Harapan site (81 spe-
cies). There were no significant differences in ant
species richness between land-use types, neither
in BDNP (ANOVA, F3, 10= 1.26, P = 0.340) nor
in Harapan (ANOVA, F3, 15 = 0.37, P = 0.779).

11 - AM7 - Agricultural land use alters species composition.indd 3 16-Jul-15 3:55:38 PM

 4 Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat, Pudjianto,
Dadan Hindayana, Yann Clough & Teja Tscharntke, Damayanti Buchori

Nevertheless, species accumulation curves show
differences in ant species diversity between the
different sites and land-use types (Fig. 2).
Sites within each land-use type had
a higher similarity of ant species composition
than sites from different land-use types (Table
2). NMDS ordination analysis showed that there
were significant differences in ant community
structure between land-use types in both, BDNP
(ANOSIM, R = 0.737, P = 0.001) and Harapan
(ANOSIM, R = 0.652, P = 0.001) sites (Fig. 3).
In both, BDNP and Harapan sites, nine
ant species were recorded in all habitat types,
i.e. forest, jungle rubber, rubber plantations and
palm oil plantations (Fig. 4). Several ant species
dominated the study plots (Fig. 5) that are mostly
categorized by Brühl & Eltz (2010) as non-forest
species and do not normally occur in forest habi-
tats, i.e. Anoplolepis gracilipes (Smith, 1857),
Dolichoderus sp. 01 and 02, Odontoponera
denticulate (Smith, 1858), Monomorium sp. 02,
Technomyrmex sp. 02, Oecophylla smaragdina
(Fabricius, 1775), Nylanderia sp. 02, and Cre-
matogaster sp. 01.
DISCUSSION
Transformation of near-primary forests to agro-
forests and plantations is often accompanied by
drastic changes in biodiversity. Against our ex-
pectation, species richness did not differ signifi-
cantly between the forest, jungle rubber, rubber
and oil palm sites. However, species composition
differed strongly between land-use types. Ant
communities in rubber and oil palm plantations,
both in the BDNP and Harapan sites, could be
clearly distinguished from forest and jungle rub-
ber communities. Forest and jungle rubber sites
were more similar, even partly overlapping in one
of the two areas studied.

Fig. 1. Study area in two sites of Bukit Duabelas and Harapan in Jambi Province, Sumatra. Gray colour indicates forest.

11 - AM7 - Agricultural land use alters species composition.indd 4 16-Jul-15 3:55:38 PM

 Agricultural land use alters species composition but not species richness of ant communities 5

Fig. 2. Species accumulation curves of ant species found four land use types within the two study sites, (a) Bukit
Duabelas National Park and (b) Harapan Forest. The dashed line indicates ant species richness from 15 sub-plots.

The absence of significant differences In contrast to species richness, ant com-

in ant species richness between forests and ag-
ricultural land-uses could be due to the fact that
the remaining dry lowland forests in the region
are not primary but secondary forests. Similarly,
most forests that were transformed into palm oil
plantations were not primary but secondary for-
est (as the forest plots in our project area are),
which had previously been used for logging, or
as agroforests (Koh & Wilcove 2008), so that the
ant species pool may already be eroded at the re-
gional level by past land-use changes. However,
as we discuss below, our results suggest that a
fairly large number of common and generalist ant
species, tolerant of, or specialized to, open land
and monoculture plantations, inhabit the man-
made habitats.
munity structure greatly differed between all land
use types, with differences most evident between
forests and agroforests on one hand, and the
monoculture plantations on the other. The direct
effects of the present habitat, such as differences
in available resources (food, shelter, potential
nesting sites), environmental conditions (temper-
ature, light), the open land phase of establishment
of monocultures, and indirect effects mediated
by a shift towards dominant, invasive species are
likely to explain these patterns. Ant communities
in BDNP oil palm plantations showed high simi-
larities among plots compared to other habitats
including oil palm in Harapan, which may be due
to the homogeneous understory vegetation in oil
palm plantations in the BDNP site.

11 - AM7 - Agricultural land use alters species composition.indd 5 16-Jul-15 3:55:38 PM

 6 Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat, Pudjianto,
Dadan Hindayana, Yann Clough & Teja Tscharntke, Damayanti Buchori

Fig. 3. Variation in ant community structure between study sites in the two study areas (a) BDNP and (b) Harapan,
in non-metric multidimensional scaling (NMDS) ordination (based on abundance data and a Bray-Curtis distance
metric). Forest sites are denoted by an F as the second letter, Jungle Rubber sites with J, Rubber sites with R and
Oil Palm sites by an O. Stress values are given for a 2 dimensional NMDS.

Fig. 4. Common ant species recorded from all land use types in (a) Bukit Duabelas and (b) Harapan area.

11 - AM7 - Agricultural land use alters species composition.indd 6 16-Jul-15 3:55:39 PM

 Agricultural land use alters species composition but not species richness of ant communities
Fig. 5. The most abundant ant species based on number of subplots collected from Bukit Duabelas and Harapan sites.
The species of ants found in all four
land-use types can be characterized as general-
ists, and are probably species that originate from
primary forest and tolerate the transformation to
plantations (Perfecto & Vandermeer 2006). Spe-
cies in the genera Crematogaster and Pheidole
were present in all four land-use types and are
often generalist species. The subfamily Myrmi-
cinae, in which the majority of ants species col-
lected are included, harbours many common ant
species that are widespread in warmer habitats,
and includes more than 900 described species
worldwide (Eguchi et al. 2006). There is often
competition between these generalist species and
species of the Dolichoderinae subfamily (Ander-
sen 2000), represented here for example by ants
of the genera Tapinoma and Technomyrmex, that
are also present in the four land-use types stud-
ied here. Ant species that were dominant in oil
palm and rubber plantation are generally tramp
species, i.e. species that benefit from habitat deg-
radation and human association (McGlynn 1999).
These include species of the genus Pheidole and
Tetramorium that are found in this study, which
can be invasive (Schultz & McGlynn 2000).
One of the species that is present in three
types of agricultural land use (jungle rubber, oil
palm and rubber plantations) but not the forest is
A. gracilipes. This species is well-known as in-
vasive species and thrives in disturbed areas, but
11 - AM7 - Agricultural land use alters species composition.indd 7
7
not forest. Brühl et al. (2003) also found that A.
gracilipes is the most common species on 70% of
all baits placed in oil palm plantations in Sabah,
Malaysia. A. gracilipes is one of the most inva-
sive species in the Indonesian cocoa plantations
and is associated with land-use systems with low
tree canopy cover and a small number of forest
ant species (Bos et al. 2008).
Overall, the most dominant ant species
are invasive non-forest ants such as A. gracilipes
and Odontoponera denticulata. In oil palm and
rubber plantations, O. denticulata replaced a spe-
cies of the same genus found in forest and jungle
rubber, Odontoponera transversa, These two re-
lated species can be used as bio-indicators, be-
cause they seem to have different adaptability and
different habit preferences, as already suggested
by a previous study, in which O. denticulata were
only found in urban areas, while O. transversa
were found only in relatively intact forests (Rizali
et al. 2008).
Forest ant species in the genera Cataula-
cus, Tetraponera and Polyrhachis were not com-
monly found in any of the plots, not even regu-
larly in the forest. This could be because it is more
difficult to sample the complete ant fauna in a for-
est because of its high microhabitat heterogeneity.
Tapinoma sp. 01 is abundant and very active in
Harapan site. When Tapinoma sp. 01 is abundant,
other ant species were unlikely to be present, even
16-Jul-15 3:55:39 PM
 8 Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat, Pudjianto,
Dadan Hindayana, Yann Clough & Teja Tscharntke, Damayanti Buchori
physically large ant species such as Camponotus
gigas and Polyrhachis spp.. In habitats where
dolichoderine species were not found, we found
many individuals of small species such as Mono-
morium and large species such as Oecophylla and
Tetraponera, suggesting that dolichoderines out-
compete species from other subfamilies.
To conclude, the conversion of forested
habitat results in severe changes in ant communi-
ties. While our study suggests this needs not be
accompanied by a decrease in species richness,
the identity of the species, the abundance of tramp
and invasive ants, and the dominance patterns are
different in agricultural habitats. The functional
consequences are not clear, but in terms of large-
scale biodiversity, our results suggests that any
further losses of forest habitat, including conver-
sion to jungle rubber, would result in a decrease
in regional biological diversity, as those species
dependent on forested habitats cannot persist in
monoculture plantations.
ACKNOWLEDGEMENT
This research was funded by Deutsche Forsc-
hungsgemeinschaft Germany (DFG) through a
Collaborative Research Centre (CRC 990 - EF-
ForTS) - Ecological and Socioeconomic Func-
tions of Tropical Lowland Rainforest Transfor-
mation Systems. We would like to thank the field
assistants and the administration staff of CRC
990 Jambi Office. We are grateful David Lohman
and a further anonymous reviewer for their com-
ments on our manuscript.
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 10 Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat, Pudjianto,
Dadan Hindayana, Yann Clough & Teja Tscharntke, Damayanti Buchori

Appendix 1

Ant species sampled in different land-use regimes from Harapan Forest and Bukit Duabelas National
Park (+ means present). aF = Forest, J = Jungle Rubber, R = Rubber plantation, O = Oil palm planta-
tion.
Bukit Duabelas
Harapan Forest
Subfamily National Park
No Land-usea Land-usea
Species F J R O F J R O
Dolichoderinae (Forel, 1878)
1 Dolichoderus sp. 01 + + +
2 Dolichoderus sp. 02 + + +
3 Iridomyrmex sp. 01 +
4 Loweriella sp. 01 + + +
5 Philidris sp. 01 + + + + + +
6 Philidris sp. 03 +
7 Philidris sp. 06 +
8 Tapinoma sp. 01 + + + + + + + +
9 Tapinoma sp. 02 + + + + + +
10 Tapinoma sp. 03 + + +
11 Tapinoma sp. 04 + +
12 Tapinoma sp. 05 +
13 Technomyrmex sp. 01 + + + + + + + +
14 Technomyrmex sp. 02 + +
15 Technomyrmex sp. 03 + +
Dorylinae (Leach, 1815)
16 Dorylus sp. 01 +
17 Dorylus sp.02 +
Formicinae (Latreille, 1809)
18 Acropyga sp. 01 + + + +
19 Anoplolepis gracilipes (Smith, 1857) + + + + + +
20 Camponotus gigas (Latreille, 1802) + + + + +
21 Camponotus sp. 02 + + + +
22 Camponotus sp. 03 + + + + + +
23 Camponotus sp. 05 + + +
24 Camponotus sp. 07 + +
25 Camponotus sp. 08 +
26 Echinopla sp. 01 + +
27 Echinopla sp. 02 +
28 Nylanderia sp. 01 + +
29 Nylanderia sp. 02

11 - AM7 - Agricultural land use alters species composition.indd 10 16-Jul-15 3:55:39 PM

 Agricultural land use alters species composition but not species richness of ant communities 11

Bukit Duabelas
Harapan Forest
Subfamily National Park
No Land-usea Land-usea
Species F J R O F J R O
30 Nylanderia sp. 03 + + +
31 Nylanderia sp. 04 +
32 Nylanderia sp. 05 + +
33 Nylanderia sp. 07 + + + +
34 Nylanderia sp. 08 +
35 Oecophylla smaragdina (Fabricius, 1775) + +
36 Paratrechina longicornis (Latreille, 1802) +
37 Plagiolepis sp. 01 + + + + +
38 Polyrhachis sp. 01 + + +
39 Polyrhachis sp. 02 + + + + + + +
40 Polyrhachis sp. 04 + + + + +
41 Polyrhachis sp. 05 + + +
42 Polyrhachis sp. 06 +
Myrmicinae
43 Acanthomyrmex sp. 01 + +
44 Acanthomyrmex sp. 02 +
45 Acanthomyrmex sp. 03 +
46 Aphaenogaster sp. 01 + + + + + + +
47 Calyptomyrmex sp. 01 +
48 Cardiocondyla sp. 01 + + + + +
49 Cardiocondyla sp. 02 + +
50 Cataulacus sp. 01 + + +
51 Crematogaster sp. 01 +
52 Crematogaster sp. 02 + + + + + +
53 Crematogaster sp. 03 + + + + + + + +
54 Crematogaster sp. 04 + + + + + + + +
55 Crematogaster sp. 05 +
56 Crematogaster sp. 14 +
57 Lophomyrmex sp. 01 + + +
58 Lophomyrmex sp. 02 + + +
59 Lordomyrma sp. 01 +
60 Lordomyrma sp. 02 +
61 Lordomyrma sp. 03 + +
62 Meranoplus sp. 01 + +
63 Monomorium floricola (Jerdon, 1851) + + +
64 Monomorium sp. 02 + + + +

11 - AM7 - Agricultural land use alters species composition.indd 11 16-Jul-15 3:55:40 PM

 12 Ratna Rubiana, Akhmad Rizali, Lisa H. Denmead, Winda Alamsari, Purnama Hidayat, Pudjianto,
Dadan Hindayana, Yann Clough & Teja Tscharntke, Damayanti Buchori

Bukit Duabelas
Harapan Forest
Subfamily National Park
No Land-usea Land-usea
Species F J R O F J R O
65 Monomorium sp. 03 + +
66 Myrmicaria sp. 01 +
67 Pheidole sp. 01 + + + + +
68 Pheidole sp. 02 + + +
69 Pheidole sp. 03 +
70 Pheidole sp. 04 + + + +
71 Pheidole sp. 05 +
72 Pheidole sp. 06 +
73 Pheidole sp. 07 +
74 Pheidole sp. 08 + +
75 Pheidole sp. 09 +
76 Pheidole sp.10 +
77 Pheidole sp. 11 + + + + + +
78 Proatta butteli (Forel, 1912) + +
79 Recurvidris sp. 01 + +
80 Recurvidris sp. 02 +
81 Solenopsis sp. 01 +
82 Solenopsis sp. 02 +
83 Strumigenys sp. 01 +
84 Tetheamyrma sp. 01 + + + + +
85 Tetramorium sp. 01 + + + + +
86 Tetramorium sp. 02 + + + +
87 Tetramorium sp. 03 +
Ponerinae
88 Anochetus sp. 01 +
89 Cryptopone sp. 01 + +
90 Diacamma rugosum (Le Guillou, 1842) + +
91 Emerypone sp. 01 +
92 Hypoponera sp. 01 + + +
93 Leptogenys sp. 01 + + + + +
94 Mesoponera sp. 01 + + + + +
95 Myopias sp. 01 +
96 Odontomachus sp. 01 + + + + + + + +
97 Odontoponera denticulata (Smith, 1858) + + + +
98 Odontoponera transversa (Smith, 1857) + +
99 Platythyrea sp. 01 +

11 - AM7 - Agricultural land use alters species composition.indd 12 16-Jul-15 3:55:40 PM

 Agricultural land use alters species composition but not species richness of ant communities 13

Bukit Duabelas
Harapan Forest
Subfamily National Park
No Land-usea Land-usea
Species F J R O F J R O
100 Platythyrea sp. 02 +
101 Ponera sp. 01 + + + +
102 Ponera sp. 02 +
Pseudomyrmecinae
103 Tetraponera sp. 01 + + + + + +
104 Tetraponera sp. 03 + +

ASIAN MYRMECOLOGY
A Journal of the International Network for the Study of Asian Ants
Communicating Editor: Martin Pfeiffer

11 - AM7 - Agricultural land use alters species composition.indd 13 16-Jul-15 3:55:40 PM

11 - AM7 - Agricultural land use alters species composition.indd 14 16-Jul-15 3:55:40 PM